A natural history notebook and project of the Missouri Botanical Garden

Costa Rica

Damaged ecosystems don’t recover overnight, but sometimes that’s all the time that they get. CCSD scientist Leighton Reid describes new research about tropical secondary forests in southern Costa Rica, including how long these young forests last, what’s at stake, and how we can keep them around longer.

Regrowing tropical forests on marginal farm lands is one of the main ways that humans can prevent runaway climate change. With ample moisture and long growing seasons, tropical trees often can grow quickly and pull large amounts of carbon out of the atmosphere, storing it in their wood and keeping it from trapping heat. At the same time, young forests provide habitat for plants and animals and improve water quality for humans, among many other benefits.

But even in a moist, tropical climate, trees don’t grow instantly. Typically, it takes many decades for a recovering forest to stock up all of the carbon that it can hold. And it can take even longer for some plants (like orchids) and animals (like antbirds) to return. If a forest starts to grow back, but then someone cuts it down again, these time-dependent benefits never accrue.

In other words, the hopes and expectations that many people have for young tropical forests depend on young tropical forests growing old. So do they? Our new study suggests not.

To find out how long secondary forests persist, I teamed up with Matthew Fagan, a landscape ecologist at the University of Maryland Baltimore County, and Rakan Zahawi, director of the Lyon Arboretum, as well as two students, James Lucas at Washington University and Joshua Slaughter at UMBC.

We studied a set of historical, aerial photos from southern Costa Rica, which covered the time period from 1947-2014. Previously, Zahawi and colleagues had classified which areas in each photo were forest and which areas were farms or other non-forest land uses. By comparing the maps they made for each year, we were able to see where and when new forests appeared and how long they remained as forest before they were converted to some other land use (mostly farms).

The young forests did not last long. Half of the new forests disappeared before they were 20-years old. And 85% were cut down before they were 54-years old. Larger forests and forests near rivers lasted longer.

First, the bad news. Twenty years is not even close to the amount of time it takes for a young forest to become as diverse as an old-growth forest. For example, vascular epiphytes like orchids and bromeliads take more than 100 years to fully recover in young forests.

Carbon storage will also take a hit. If forests elsewhere in Latin America are as ephemeral as forests in southern Costa Rica, then carbon stocking over the next thirty years may be reduced by an order of magnitude.

Ephemeral forests could just be a problem in Costa Rica, but another study shows that secondary forests in eastern Peru have even shorter lifespans. There, secondary forests are cleared at a rate of 3-23% per year. Compared to that, the 2-3% per year rate of loss in southern Costa Rica is considerably better. And that’s not a good thing. Clearly we need more research on secondary forest persistence from other places.

There is some good news, though. Even though many new forests were short-lived, the ones that survived were predictable. And if we can predict where new forests will survive, we should also be able to help them survive longer. Larger forests and forests close to rivers were cut down less often than small forests and forests far from rivers. This suggests that restoring large, riparian forests could be a smart investment.

Governments and other organizations can also help forests persist by creating incentives for long-term forest management, providing resources to enable long-term management, and ensuring that local people will be able to enjoy the benefits that old forests provide.

We hope that this work will lead to stronger restoration commitments. Right now, dozens of countries are setting big targets for forest restoration. For example, in 2012 Costa Rica committed to restore a million hectares of degraded land by 2020 (an area about one fifth the size of the country). There is a great opportunity for Costa Rica and other ambitious countries to plan for long-term forest restoration.

If we can begin to restore a million hectares of forest by 2020, why not plan to restore a million hectares of 100-year old forest by 2120?

A trail through secondary forest at the Las Cruces Biological Station in southern Costa Rica. Photo by J. Leighton Reid.

Sometimes farmlands quickly regrow tropical forests on their own, but other times they don’t. Dr. Karen Holl, a professor at the University of California Santa Cruz, gives some rules of thumb for when we can save money on tropical forest restoration by letting nature do the work, and when we may need to invest in tree planting.

Ambitious targets are being set to restore tropical forest because of their importance in storing carbon, regulating water cycles, conserving biodiversity, and supporting the wellbeing of people who live in tropical countries. For example, the 20 × 20 Initiative aims to restore 20 million hectares of tropical forest in Latin America by 2020. This represents an area slightly smaller than the country of Ecuador. One big question is: How are we going to restore forests at this scale with limited funds?

One of the cheapest ways to restore forest is to let nature do the work and leave forests to recover on their own. This works in some sites where forests regenerate quickly. In other cases, usually sites that have been used intensively for agriculture, the land may be covered by tall grasses (up to 3 meters, or 10 feet high) for years. Our past research shows that even within a small region, the rate of natural forest recovery varies greatly.

Natural forest recovery is highly variable in southern Costa Rica, even after a decade of recovery. Left: slow recovery on a former farm, still dominated by non-native grasses, with an open canopy and little tree recruitment. Right: speedy recovery on a former farm, with virtually no grass cover, a closed canopy, and diverse tree recruitment. Photos by Andy Kulikowski.

So, how do we predict which sites will recover quickly and which ones need some help in the form of clearing pasture grasses and planting trees? If we could develop some rules of thumb it would help land managers to more efficiently allocate scarce restoration funds.

To answer this question, we drew on our long-term study on tropical forest restoration in southern Costa Rica. We have research plots at 13 different sites where we removed the land from agriculture and let the forest recover on its own. Each year we measure grass cover, tree canopy cover, and how many and what species of new tree seedling establish in the plots. We have also quantified the forest cover surrounding the plots, the nutrients in the soil, and how long cows had grazed the sites in the past.

We found that two easy-to-measure variables explained on average two-thirds of variation in forest recovery 7 years later; those were the amount of grass cover and tree canopy cover measured after only 1.5 years. Plots that had more canopy cover and lower grass cover early on had a closed tree canopy and lots of forest tree seedlings from many species after nearly a decade. We were surprised that the amount of surrounding forest cover and soil nutrients did not explain much of the variation in forest recovery.

Rules of thumb for predicting tropical forest regeneration on farmlands. Forests grow back quicker when there is not too much grass, a little bit of shade, and many tree seedlings already present. Illustrations by Michelle Pastor.

Of course, our results need to be tested in other recovering tropical forests. But, if they hold true, this is good news! It means that land owners and managers just need to wait a year or two and then measure the tree canopy and grass cover. If some trees have established and are starting to shade out the grasses, land managers can use the low cost method of leaving the site to recover naturally. If the site is mostly a monoculture of dense grass, then the site is a good candidate to plant native trees. Planting trees takes more resources since it is necessary to clear around the native tree seedlings for a couple of years until they grow taller than the grasses. At least now there are some general guidelines to help chose where to invest the extra effort.

CCSD scientist Leighton Reid and Lyon Arboretum director Rakan Zahawi write about giant fig tree cuttings: how to make them and why some grow better than others.

Choosing the right species to include in a restoration project is a hard choice, but in the economy of nature, some species earn a bigger ROI than others. For example, Pacific sea otters maintain kelp forests by eating sea urchins, and wolves in Yellowstone National Park allow aspen groves to regenerate by scaring away tree-munching elk. These vital creatures are called “keystone species” because they hold ecosystems together, much like the keystone in an arch.

Plants can be keystone species too. Around the world there are about 800 species of fig trees, and they hold tropical forests together by providing food for a wide array of animals. On any given day, the busiest tree in a rain forest is likely to be a fig tree with fruits. Monkeys, birds, bats, and others gather at fig trees to eat, and in the process, they deposit seeds of other plant species that they have been carrying in their guts. This chain of events, repeated day after day, often turns the area beneath a fig tree into a hotspot of plant diversity.

A few years ago, we had an idea to plant keystone fig trees in young forests in Costa Rica. We wanted the figs to grow as fast as they could, so instead of planting seedlings, we planted cuttings – big ones. With help from our local collaborator, Juan Abel Rosales, we cut dozens of twelve foot-long branches from eight species of fig trees. We stripped off all of their leaves to keep them from drying out, and then we planted our figs trees in shallow holes.

Rakan Zahawi (delighted!) poses with a three year-old fig stake.

To our delight, many of the fig trees grew!

The ones that did the best came from a special group, the subgenus Urostigma. Many figs in this group have a unique life strategy. They begin their lives in the top of a tree when their tiny seeds are deposited on a branch by a bird or some other animal. As they grow in the treetop, they send long roots down to the ground, and these roots harden and fuse together, forming a lattice-like trunk. Over time, these figs kill their host trees by taking most of the water, nutrients, and light. They also keep the host tree from growing outwards, giving them the nickname “strangler figs”. Maybe the ability to transform a flimsy, dangling root into a solid trunk is related to these figs being able to grow from cuttings.

To find out how well our planted fig cuttings might survive over the long-term, we also tracked down some fig cuttings that we had planted in 2004. We were happy to learn that out of the trees that survived for their first three years of life, all of them were still thriving a decade later.

Full disclosure: planting large cuttings is not a new idea. Farmers in many parts of the tropics plant trees this way to create ‘living fences’ – with all of the normal fixings like gates and barbed wire, but with a row of living trees instead of dead posts. The advantages for farmers are many – their fences don’t rot and fall apart (that happens quickly in the tropics); the trees provide shade for cattle; they have a constant source of new fence posts (by cutting off a limb); and in some cases they can feed the young shoots to livestock.

Big cuttings have big benefits for restoration too. Not only are planted trees already several feet tall, you also get to skip the pricey nursery phase, and, most excitingly, cuttings have a tendency to fruit quickly.

Some of our young fig trees are now making fruit, but we will have to wait a bit longer to see whether they start attracting more big animals and whether those animals carry more tree seeds into our young forests. For now, we can say that others who are interested in growing keystone figs for forest restoration may have the best luck by working with the stranglers.

How to grow an instant fig tree. (A) Remove a long, thin branch segment from an adult tree. The red arrow shows a cut branch. (B) Strip the cuttings of their leaves to keep them from drying out, then carefully transport cuttings so as not to damage cortical tissue. Here, cuttings are padded by a foam mattress. (C) Remove the bark from a ring on the cutting to promote root growth. Here, a ring is being cut about 20 cm (8 in) above the base so that it will be just below the soil surface when planted. (D) Dig a shallow hole and plant the cutting. Be sure that the cutting is firmly planted to prevent it from toppling, but take care not to compact the soil too much around its roots. Photos by Rakan Zahawi.

During a recent trip to Costa Rica, CCSD scientist Leighton Reid toured La Sabana, Costa Rica’s largest urban park, with Wilmar Ovares, an instructor at the Universidad Estadal a Distancia who has been studying the recovery of bird diversity following large-scale replacement of exotic trees with native ones.

The first time I visited La Sabana Metropolitan Park in downtown San Jose was in 2005. At that time it was essentially a eucalyptus woodland; the tall trees with peeling bark stretched upwards above the soccer fields and hiking trails. No longer. La Sabana has gotten a makeover in the last few years – and for the better.

The old La Sabana (exotic eucalyptus trees in background – slated for removal in the near future) and the new (native trees and a eucalyptus stump in the foreground). The tree at right (unidentified species; Solanaceae) is a natural recruit, dispersed to the site by a bird or a bat. The slightly curved tree at center-right has died, but it can still serve as a bird perch and might facilitate the dispersal and establishment of a new, native tree in its place.

The project was initiated as a collaboration between three institutions: the National Biodiversity Institute (INBio), the National Sports and Recreation Institute (ICODER), and Scotiabank. Beginning in 2010, this collaborative removed most of the exotic trees in La Sabana and replaced them with more than 5000 native trees, representing 234 species. Not all of the species are native to the central valley, but all are native to the country.

Although the planted trees are still quite small, one short-term indicator of project success is the recovery of bird diversity in the park. On my first visits to La Sabana prior to 2010, the birding was slow, with occasional excitement when I would stumble on a eucalypt in flower – abuzz with warblers, orioles, and hummingbirds. Now, Wilmar Ovares finds that the number of bird species has increased by more than a third. Many of the new species are migrants, which breed in North America and winter in Central America. Others have drifted in from nearby riparian forests along the Rio Torres and the Rio Maria Aguilar. In some cases, birds and other animals have carried in and deposited native tree seeds, complementing the plantings.

This Annona cherimola (Annonaceae) recruited on its own at this site in La Sabana. Its large seed may have been dispersed by a bird, a squirrel, a raccoon, or even a human. The source of the seed may have been the Rio Torres, which flows through a riparian forest not far from the park. This tree species is a conservation priority species in the Central Valley.

To be clear, this project is not strictly ecological restoration; below the new trees is a manicured lawn, and it is likely to remain that way for some time. The work is better classified as rehabilitation – a return of some elements of the local biodiversity, but by no means all of it. This approach is sensible given that the park is a resource for human recreation – not just a habitat for plants and animals. Still, if the park administration wished to go further, they could consider introducing some native, understory shrubs, ground-layer plants, and epiphytes, all of which would enhance bird diversity and enrich the experience of visitors.

Acerola (Malpighia glabra, Malpighiaceae) in a recent planting in La Sabana. This tree/shrub produces edible fruits, which are not very sweet. (Thanks to Amy Pool for correcting a previous misidentification!)

The last vestiges of the old La Sabana. This area near the stadium is dominated by eucalyptus (from Australia) and Cupressus lusitanica (from Mexico and northern Central America). Wilmar Ovares finds a lower diversity of bird species in these exotic tree groves, despite their much greater stature. Though it cannot be denied that when the eucalyptus are flowering, warblers, orioles, hummingbirds and others may be found in large numbers feeding on the nectar and nectar-eating insects.

Wilmar Ovares has been monitoring changes in the bird community in La Sabana as a result of the native tree rehabilitation.

Estefania Fernandez is a Bascom Fellow who recently finished her master’s thesis at the University of Montpelier, France. Last year, Estefania wrote about her preliminary results on tropical forest restoration and vascular epiphyte reintroductions in Costa Rica. Here, she describes the final results, recently published in Restoration Ecology.

A transplanted bromeliad, Aechmea dactylina flowering in a 10-year old tree plantation.

Vascular epiphytes are plants that germinate and root on other plants without taking their nourishment from their host plant, and they represent 50% of the flora in some tropical forests and 9% of all vascular plants worldwide. If you are a plant lover, then you most likely have one or several vascular epiphytes in your house. Some of the most appreciated horticultural families include orchids (Orchidaceae), aroids (Araceae), and bromeliads (Bromeliaceae).

Vascular epiphytes also play key roles in our ecosystems. They are crucial to forest water and mineral recycling as they intercept rainfall and prevent rapid run-off and nutrient leaching. Vascular epiphytes are also exceptional microhabitats where invertebrate communities find refugia and birds and arboreal mammals forage.

Transplanted individual of Werauhia gladioliflora

Despite their importance in forest ecosystems, vascular epiphytes are rarely taken into account in forest restoration. This is problematic because vascular epiphytes are often among the slowest plants to recolonize regenerating forests.

In 2015-2016, I tested whether transplanting epiphytes into young restoration sites could be a viable strategy to accelerate their reestablishment. I used a bromeliad for my experiment, Werauhia gladioliflora (H. Wendl.) J.R. Grant, which was common in remnant forest but had not been found during epiphyte surveys in nearby restoration areas. In March-June 2015, I transplanted 60 bromeliads into three restoration plantations near Las Cruces Biological Station in southern Costa Rica. I revisited the sites in January-February 2016, nine months after transplantation, to monitor survival and arthropod recolonization.

Happily, over 75% bromeliads survived and the number of arthropods on branches with bromeliads was seven times greater than in branches without bromeliads. Additionally, I observed that bromeliads buffered the local microclimate; during the driest and hottest times of the day, the interior of the bromeliads was moister and cooler than ambient air.

Transplanted individuals of Werauhia gladioliflora (left) hosted considerably more arthropods in their rosettes than could be found on the stems of trees that had not received a transplant. GN, JG, and MM are three study sites near Las Cruces Biological Station in southern Costa Rica. Photo by Dave Janas.

Restoring arboreal refugia

My research suggests that transplanting fallen epiphytes onto trees in restored sites contributes to the recovery of vascular epiphyte diversity in these ecosystems and has the additional benefits of bringing back arthropod diversity to these sites. Epiphytes, and specifically “tank” epiphytes that retain water in their rosettes, help stabilize microclimatic conditions, a critical function in light of climate change, which may put arboreal communities at special risk. Indeed, the body temperature of many animals such as invertebrates entirely depends on ambient temperatures but rising temperatures could push arboreal animal communities to the ground. Epiphytes offer ideal refugia from high temperatures and drought and their presence in tree canopies and understory is critical to preserve arboreal animal communities. Transplanting other epiphyte families or even entire epiphyte communities found on fallen branches could be tested in the future to broaden this strategy.

Estefania inspects a flowering individual of an Aechmea dactylina transplant

This work was supported by a grant from the National Science Foundation.

Planting tree islands has many of the benefits of larger plantations, but entails significantly less cost. Karen Holl (University of California, Santa Cruz), Leighton Reid (Missouri Botanical Garden), and Zak Zahawi (American University of Beirut) describe recent findings on tree seedling recruitment in a long-term experiment in southern Costa Rica.

Over the past few years there have been a growing number of commitments at the global, national and regional scale to restore forests because of their importance to conserve biodiversity, sequester carbon, reduce erosion, and provide goods and services to people. For example, Initiative 20×20, led by the International Union for the Conservation of Nature, aims to restore 20 million hectares of tropical forest by 2020, an area roughly equivalent to the size of Uruguay or Nebraska.

A common strategy to restore forests is to plant trees. But, the big question is: where will the money come from to plant billions of trees when there are so many pressing needs? As restoration ecologists, we started thinking about how we could most efficiently allocate resources to get the best bang for the buck and restore the largest area of forest.

Trade-offs in forest restoration strategies. Planting fewer trees leaves more to chance and can require more time, but tree plantations are more expensive and leave a bigger ecological footprint. Our study tests an intermediate option, and after 10 years it appears to provide a good balance. Figure modified from Corbin & Holl (2012).

Starting over 10 years ago, we set up a large-scale tropical forest restoration experiment in southern Costa Rica to test two ideas.

First, we tried planting tree “islands”. The idea is to plant groups of trees that attract birds and bats, which disperse most tropical forest tree seeds. The tree canopy also shades out light-demanding grasses that can outcompete tree seedlings. In one experimental treatment, we planted tree islands that covered about 20% of 50 × 50 m plot of former cattle pasture. We compared that to plots where no trees were planted (natural recovery) and to the more intensive (and more typical) restoration strategy of planting trees in rows throughout the plot (plantation).

Second, we asked: is it only possible to restore forest near remnant forests or can you restore forest anywhere in the landscape? This is important information to help guide forest restoration efforts. To do this we set up our entire experiment at 13 sites, some of which were mostly surrounded by agricultural land and some of which were adjacent to the largest remaining forests in the region.

Then we monitored establishment of new tree seedlings in our research plots over a decade. We compared the number of seedlings, number of species, and types of species in the restoration plots with those found in the nearby forest to evaluate how well the forest is recovering.

The tree island planting method not only saves money on buying, planting, and maintaining seedlings, but it also results in a more heterogeneous distribution of trees, so it looks more like a natural forest.

Profuse tree seedling and sapling recruitment in the understory between two tree islands in southern Costa Rica.

We counted over 6000 tree seedlings, 88% of which have seeds that are dispersed by animals. On average there were many more tree seedlings in the tree island and plantation treatments than in the natural recovery plots. These results suggests that some tree planting helps the forest to recover faster, but that it is not necessary to plant the whole area with trees. The tree island planting method not only saves money on buying, planting, and maintaining seedlings, but it also results in a more heterogeneous distribution of trees, so it looks more like a natural forest.

Even though there were many tree seedlings in the island and plantation plots, on average there were less seedlings of tree species that have big seeds (>0.5 cm/0.2 inches across) compared to mature, reference forests. It seems that the larger-seeded species that are common in mature forests are much slower to colonize restored sites, likely because they are eaten and dispersed by a small number of larger animals, such trogons and agoutis. Many of those dispersers are less likely to visit early successional forest.

Small frugivores, small seeds. Most of the birds we see in these experimental plots are small-gaped omnivores (e.g., Yellow-bellied Elaenia, Elaenia flavogaster, left), but it usually takes large-gaped species to disperse larger seeds1. The figure at right shows the maximum fruit size that a bird species with a given gape size was able to consume in a cloud forest in central Costa Rica (modified from Wheelwright (1985)). In our experiment, small seeds were ubiquitous, but large seeds were mostly absent.

We were surprised that the amount of forest cover around the experimental plots had a weak effect on the number of seedlings establishing. In other words, isolated plots had just as many tree seedlings as plots right next to old-growth forests. We think that this is likely due to the fact that there are many trees in the agricultural landscape surrounding our plots; these trees include remnant trees, living fence rows, and riparian corridors. Trees in the landscape can serve an important role in both providing sources of seeds and stepping stones for the movement of seed-dispersing fauna. We anticipate that having forest nearby will be more important in future years as these forests build up greater diversity of rare, large-seeded species. Nonetheless, our results suggest that there are good prospects for restoring forests in many locations in this landscape.

Our key finding is that planting tree islands can be a cost-effective way to restore tropical forests at our study site in Costa Rica, but we hasten to note that the strategy should be tested in other locations, particularly areas with fewer forest elements in the surrounding countryside. Our study also demonstrates that tropical forests can recover some species quickly but it will take many decades, if ever, for forests to fully recover. So, preserving existing rain forests is critical to conserve biodiversity and the services they provide to people.

Diverse tree cover in an agricultural landscape in southern Costa Rica. Remnant trees in pastures, trees along fence rows, and riparian forests provide important sources of flora and fauna to speed up forest recovery.

1See Melo et al. (2009) for an example to the contrary: small-gaped animals dispersing fairly large fruits and seeds.

This work was supported by a grant from the National Science Foundation.

Andres Santana is the graduate program coordinator at the Organization for Tropical Studies. During a recent fieldtrip in southern Costa Rica, he and CCSD restoration ecologist Leighton Reid compared notes on using fig stakes for ecological restoration.

Tropical beaches are many things to many people. To plants, beaches are hot, sandy, and salty – complicating their restoration.

Costa Rica has 1228 km (763 mi) of coast line – including 1016 km on the Pacific side and 212 km on the Caribbean. Along Costa Rica’s northern Pacific coast, the beach forms the natural edge of the dry forest. Farther south the adjacent forest is more humid. Giant trees, 40 m or more in height, grow right up to the high tide mark, particularly along the Caribbean.

But as with so many tropical ecosystems, Costa Rica’s coastal forests have been subject to human impacts. Many shoreline forests were cleared for cattle ranching, and exotic grasses were introduced as forage. Some of these grasses are fierce competitors and prevent tree seedlings from establishing, even long after the pastures have been abandoned.

Playa Hermosa, before (left) and after (right) planting 2-m long cuttings of a coastal fig species (Ficus goldmannii).

In 2009, a small non-profit organization, Costas Verdes, was formed to restore coastal forests along degraded shorelines, particularly wildlife refuges. The restoration work was initially challenging; tree seedlings were hard to establish along the coast because of the harsh environment – high temperatures and salinity and lack of freshwater were among the most significant obstacles. Not to mention the invasive cattle forage grasses.

Coastal restoration at Playa Hermosa

Playa Hermosa, a surfing destination on the Central Pacific coast, was among the most heavily deforested project sites. This area, part of a wetland and river estuary, was declared a national wildlife refuge in 1998. By 2009, very little forest had naturally regenerated. This led Costas Verdes to implement a restoration project at this beach. Planting plots were established where invasive grass was removed. In other areas, grasses left intact, as a comparison. It quickly became evident that tree seedlings were outcompeted by the grass. Those in the cleared plots grew better, but they still faced the other coastal habitat challenges.

Some native trees are resistant to hot substrates and high salinity, but these species were not available in tree nurseries, most of which focused on ornamental species. This meant that seedlings needed to come from locally collected and germinated seeds. We realized that this would take time to get going. Tree seedlings under 50 cm rarely survive, even if they have the proper coastal adaptations.

To accelerate the restoration, we decided to use tree cuttings rather than growing seedlings from seed. A colleague suggested Ficus goldmannii as a candidate species, so in 2011 we conducted a planting trial. We planted 225 2-m long cuttings. Of these, 195 (87%) survived their first year. By the second year all 195 survivors had become established and were quickly providing canopy cover and lowering the temperature of the sand.

An established fig stake with a dense canopy. Note the weak, patchy grass below it.

Once fig stakes created some canopy cover, we brought in other tree species – mostly from the coastal tree nursery that we created. Shade from the fig canopy also began to inhibit the invasive grasses, which require high sunlight to photosynthesize efficiently. Reduced competition with these grasses allowed other tree seedling species to survive.

In this instance Ficus cuttings turned out to be useful in promoting restoration. We have since used cuttings for other plots with similar success.